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GENETIC LOAD

In population genetics, 'genetic load' or 'genetic burden' is a measure of the cost of lost alleles due to selection ('selectional load') or mutation ('mutational load'). It is a value in the range 0 < {L} < 1, where 0 represents no load.
The concept was first formulated in 1937 by JBS Haldane, independently formulated, named and applied to humans in 1950 by H. J. Muller[1], and elaborated further by Haldane in 1957.[2]

Contents
Definition
Mathematics
Causes of genetic load
Mutational load
Selectional load
Segregational load
Creationist criticism
References
External links

Definition


Genetic load is the reduction in selective value for a population compared to what the population would have if all individuals had the most favored genotype.[3] It is normally stated in terms of fitness as the reduction in the mean fitness for a population compared to the maximum fitness.

Mathematics


Consider a single gene locus with the alleles mathbf{A} _1 dots mathbf{A} _n, which have the fitnesses w_1 dots w_n and the allele frequencies p_1 dots p_n respectively. Ignoring frequency-dependent selection, then genetic load (L) may be calculated as:
:L = {{w_max - ar w}over w_max}~~~~~~~~~~(1)
where w_max is the maximum value of the fitnesses w_1 dots w_n and ar w is mean fitness which is calculated as the mean of all the fitnesses weighted by their corresponding allele frequency:
:ar w = {sum_{i=1}^n {p_i w_i}} ~~~~~~~~~~(2)
where the i^mathrm{th} allele is mathbf{A}_i and has the fitness and frequency w_i and p_i respectively.
When the w_max = 1, then (1) simplifies to
:L = 1 - ar w. ~~~~~~~~~~(3)

Causes of genetic load


Load may be caused by selection and mutation.
Mutational load

Load caused by mutations is known as mutational load.
Selectional load

Selection occurs when the fitnesses of particular alleles are inequal, hence selection always exerts a load.
With directional selection, the allele frequencies will tend towards an equilibrium position with the fittest allele reaching a frequency in mutation-selection balance. As mutations are rare, this is effectively fixation. Consider two alleles mathbf{A}_1 and mathbf{A}_2. If w_1 > w_2, then at equilibrium, p_1 pprox 1 and p_2 pprox 0, hence ar{w} pprox w_max, and L pprox 0.
If the mean fitness is 0, the load is equal to 1, but the population goes extinct.
Segregational load

In contrast to directional selection, heterozygote advantage always exerts a load at equilibrium.

Creationist criticism


Some creationists (such as Henry M. Morris) have suggested that mutational load would increase over time and thus make populations inviable. However, they ignore the effect of selectional load acting to weed out (decrease frequency of) deleterious mutations.

References


1.
2. The cost of natural selection, JBS Haldane, , , Journal of Genetics, 1957
3. Some possibilities for measuring selection intensities in man, JF Crow, , , Hum. Biol, 1958

External links



''The Cost of Natural Selection Revisited'', Leonard Nunney, Ann. Zool. Fennici 40:185-194 (pdf file)

Genetic load, from ''Evolution A-Z'' by Mark Ridley

"Understanding Genetic Load" - a paper by Dr. L. Monroe

Creationist Claim CB120: Genetic load from ''An Index to Creationist Claims'' by Mark Isaak

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